Cytotaxonomy of Lychnophora Mart. (Asteraceae: Vernonieae: Lychnophorinae) Species

CARYOLOGIA Vol. 60, no. 1-2: 21-28, 2007

Cytotaxonomy of Lychnophora Mart. (Asteraceae: Vernonieae: Lychnophorinae) species

Mansanares1 Mariana Esteves, Eliana Regina Forni Martins* and Joa˜o Semir

Departamento de Botaˆnica, Instituto de Biologia, Universidade Estadual de Campinas; Caixa Postal 6109, Campi- nas, SP, 13083-970, Brasil; fax:++55-19-37886168. 1 Post-graduate Student of Biologia Vegetal Pos-graduate Course, UNICAMP.

Abstract — Most species of the genus Lychnophora Mart. are endemic to the Brazilian ”campos rupestres”ofMinas Gerais, Bahia and Goia´s, with high degree of endemism in many species. There is disagreement between different authors regarding delimitation of the species and, consequently, the amount of species (from 11 to 68). This inter- pretation difference leads to sinonimization and the transference of several species to closely related genera. The cytotaxonomic study of species of Lychnophora was made aiming at increase of knowledge of chromosome character- istics that could be useful to the understanding of the taxonomy of the group as a whole. Chromosome numbers of about eighteen species were determined, with 2n=34, 36 or 38. These chromosome numbers were distributed among species of four sections of Lychnophora,sothey can’t be used as distinctive characters for intergeneric and infrage- neric levels below section level. However, chromosome numbers were very important for the differentiation of some species of Lychnophora, whose taxonomic limits have been questioned. Other karyotype characters were analysed in three species of the subtribe, like chromosome size and morphology, showing constancy of these characters. The chromosomes are small, with 1.0 to 2.58µm, and they are mainly metacentric, however some submetacentrics were observed.

Key words: Asteraceae, chromosome, cytotaxonomy, karyotype, Lychnophora, Lychnophorinae, Vernonieae.

INTRODUCTION According to Semir (1991), the genus Lychno- phora is divided in 6 sections: Lychnophora, Lych- The genus Lychnophora was described by nophoriopsis, Lychnophorioides, Lychnocephaliop- Martius (1822), who considered the presence of sis, Sphaeranthus and Chronopappus. The author glomerular inflorescences and achenes with de- classified these sections based on inflorescence ciduous pappus as differentiating features. It oc- morphology and presence or absence of sheath or curs exclusively in Brazilian’s “campos rupestres”, petioles. with a great number of endemic species. Lychno- Before 1980, chromosome numbers for Lych- phora shows some taxonomic problems related to nophora sensu Semir (1991) species were not the species circunscription and delimitation, with known. Even in present day karyotype informa- acceptance of 11 (Coile and Jones 1981), 34 tion, as chromosome length and morphology, is (Robinson 1999), or 68 species (Semir 1991). not available. The initial suggested chromosome Coile and Jones (1981) and Semir (1991) number was n=17/2n=34, for L. ericoides, L. to- pointed out the possible occurrence of natural in- mentosa, L. heterotheca (= L. candelabrum) and L. terspecific hybrids as one of the factors that com- diamantinana (Coile and Jones 1981). Later, plicated the genus’ taxonomy. Also, there is some Jones (1982) cited 2n=36 for Eremanthus reflexo- conflict in circumscription of the genera in sub- auriculatus (= L. reflexo-autriculata) and Carr et tribe Lychnophorinae (Coile and Jones 1981; al. (1999) reported an atypical chromosome MacLeish 1984; 1987; MacLeish and Schu- number (2n=18 +1B) for L. phylicifolia. maker 1984; Semir 1991; Robinson 1992; 1996a; Recently, Mansanares et al. (2002) carried b; 1999; Hind 1995 2000). out cytotaxonomic studies in Lychnophora Mart. sensu Semir (1991) and demonstrated how those approaches are important as a subsidy for the genus taxonomy. The observed data shows variation * Corresponding author: e-mail [email protected] in chromosome numbers with 2n=34, 2n=36 and 22 mansanares, forni martins and semir

2n=38, but the most common number among though some of the species (L. angelae, L. cryp- studied species is 2n=34. According to tomerioides, L. grazielae, L. itacambirensis, L. mu- Mansanares et al. (2002), L. ericoides and L. gard- tica, L. nanuzae, L. prostrata and L. sobolifera) neri (sensu Semir 1991) show different chromo- have not yet been published and were only men- some numbers (2n=34 and 2n=36, respectively), tioned in Semir’s doctorate dissertation. reinforcing the species segregation, in opposition The mitotic root-tips were pre-treated by 5 to Coile and Jones (1981), who considered them hours in 8-hydroxiquinoline 0,002M solution, at only as L. ericoides. 15°C, fixed in Carnoy solution (acetic acid − etha- The present study, aimed at enlarging the cyto- nol (1:3) and then, stained according to HCl/ taxonomic knowledge of Lychnophora, presents Giemsa technique (Guerra 1983). chromosome numbers for other species and For the meiotic study, flower buds were also karyotypes of some of them. The results were fixed in Farmer’s solution and stored in ethanol compared with the different taxonomic treat- 70 at 4°C. Squashes of pollen mother cells were ments for the genus by Coile and Jones (1981), made using the acetocarmine 1,2% technique Semir (1991) and Robinson (1999). (Medina and Conagin 1964). The chromosome number counts were based on at least 20 metaphase plates from different individuals from each species, except L. villosissima MATERIAL AND METHODS (Population 01), for wich 59 metaphase mitotic and 61 meiotic cells were observed (Table 1). Eighteen Lychnophora species were analysed Ideograms were prepared for three species: (Table 1), all collected in “campos rupestres” of chromosome length was measured in at least 10 several areas of the State of Minas Gerais, as Dia- metaphase plates from each species/population, mantina, Montes Claros and Gra˜o Mogol, and of using the average measure for chromosome, of the State of Bahia, in Rio de Contas. Voucher chromosome length and centromeric position. specimens were deposited at the herbarium of Chromosome nomenclatural morphology Campinas State University (UEC). adopted here was suggested by Guerra (1986). Lychnophora species were analysed and identi- For karyotype characterization measures as TCL fied according to Semir's (1991) account, al- (total chromosome length), CI (centromeric in-

Table 1 — Species and populations of Lychnophora (subtribe Lychnophorinae) and respective localities and vouchers.

Species Local Population Voucher Lychnophora sect. Lychnophora L cryptomerioides Semir and Leitaõ Filho (ine´d.) MG, Diamantina Pop01 Mansanares et al. 163 L. granmogolensis (A.P.Duarte) Semir and Leitaõ Filho (ine´d.) BA, Rio de Contas Pop01 Moraes and Aona MDM452 L. itacambirensis Semir and Leitaõ Filho (ine´d.) MG, Itacambira Pop01 Mansanares et al. 266 L. martiana Gard. MG, Juramento Pop01 Mansanares et al. 180 MG, Montes Claros Pop02 Semir and Duthil s/n° L. mutica Semir and Leitaõ Filho (ine´d.) MG, Santana do Riacho Pop01 Mansanares and Kinoshita 270 L.nanuzae Semir and Leitaõ Filho (ine´d.) MG, Diamantina Pop01 Mansanares et al. 143 L. prostrata Semir and Leitaõ Filho (ine´d.) MG, Diamantina Pop01 Mansanares et al. 00/26 L. ramosissima Semir and Leitaõ Filho (ine´d.) MG, Diamantina Pop01 Mansanares et al. 171 L. rosmarinifolia Mart. MG, Diamantina Pop01 Mansanares et al. 00/29 MG, Diamantina Pop02 Mansanares et al. 00/36 L. salicifolia Mart. MG, Juramento Pop01 Mansanares et al. 191 L.sobolifera Semir and Leitaõ Filho (ine´d.) MG, Diamantina Pop01 Mansanares et al. 110 L. villosissima Mart. MG, Diamantina Pop01 Mansanares et al. 00/23 MG, Diamantina Pop02 Mansanares et al. 00/37 MG, Diamantina Pop03 Mansanares et al. 00/93 L. uniflora Schultz-Bip. BA, Rio de Contas Moraes and Aona MDM 508 Lychnophora sect. Lychnophorioides L. angelae Semir and Leitaõ Filho (ine´d.) MG, Serra do Cipo´ Pop01 Mansanares and Verola 377 L. leucodendron (Mattf.) Semir and Leitaõ Filho (ine´d.) BA, Rio de Contas Pop01 Moraes and Aona MDM 503 BA, Rio de Contas Pop02 Moraes and Aona MDM521 L. sincephala Gard. MG, Diamantina Pop01 Mansanares et al.232 Lychnophora sect. Lychnocephaliopsis L. grazielae Semir and Leitaõ Filho (ine´d.) MG, Serra do Cipo´ Pop01 Mansanares and Verola376 Lychnophora sect. Chronopappus L. markgravii G.M. Barroso MG, Graõ Mogol Pop01 Aona 701 cytotaxonomy of lychnophora 23 dex) and TF% (asymmetric index) were used RESULTS (Huziwara 1968). Cells with well spread chromosomes were ob- Three different chromosome numbers were served through a common photomicroscope. obtained in eighteen species of Lychnophora Photographs were taken using Agfa Pan ISO 25 (sensu Semir 1991), 2n=34, 2n=36 and 2n=38 film. (Table 2, Figures 2 − 3). All these chromosome

Table2—Chromosome numbers in species of Lychnophora, as found in literature for Lycnhophorinae subtribe and present data (*: chromosome counts reported for the first time).

Species n 2n Reference Lychnophora sect. Lychnophora L. cryptomerioides Semir and Leitaõ (ine´d.)* 18 36 Present work L. diamantinana Coile and Jones 17 - Coile and Jones, 1981 17 34 Mansanares et al. 2002 L. ericoides Mart. 17 - Coile and Jones, 1981 -34Mansanares et al. 2002 L. gardneri Sch. Bip. - 36 Mansanares et al. 2002 L. granmogolensis (A.P.Duarte) Semir and Leitaõ (ine´d.) 18+B Carr et al., 1999 34 Present work L. itacambirensis Semir and Leitaõ (ine´d.)* 18 36 Present work L. martiana Gard.* - 36 Present work L. mutica Semir and Leitaõ (ine´d.) ca.19 - Mansanares et al. 2002 -38 Present work L nanuzae Semir and Leitaõ (ine´d.)* - 36 Present work L. passerina (Mart. ex DC.) Gardner 17 34 Mansanares et al. 2002 L. pinaster Mart. 17 - Mansanares et al. 2002 L. pohlii Sch. Bip. 18 - Mansanares et al. 2002 L. prostrata Semir and Leitaõ (ine´d.) 17 - Mansanares et al. 2002 -34 Present work L. pseudovillosissima Semir and Leitaõ (ine´d.) - 38 Mansanares et al. 2002 L. ramosissima Mart.* 18 36 Present work L. rosmarinifolia Mart.* - 36 Present work L. rupestris Semir and Leitaõ (ine´d.) 17 34 Mansanares et al. 2002 L. salicifolia Mart.* 18 36 Mansanares et al. 2002 -36 Present work L. sobolifera Semir and Leitaõ Filho (ine´d.)* 18 - Present work L staavioides Mart. 34 34 Mansanares et al. 2002 L. villosissima Mart.* 18 36 Present work L. uniflora Schultz-Bip.* - 36 Present work Lychnophora sect. Lychnophorioides L. angelae Semir and Leitaõ Filho (ine´d.)* 17 - Present work L. leucodendron (Mattf.) Semir and Leitaõ (ine´d.)* - 34 Present work L. sincephala (Sch. Bip.) Sch. Bip.*-34 Present work Lychnophora sect. Chronopappus L. markgravii G.M. Barroso* 38 Present work Lychnophora sect. Lychnocephaliopsis L. cipoensis Semir and Leitaõ (ine´d.) 19 38 Mansanares et al. 2002 L. grazielae Semir and Leitaõ Filho (ine´d.)* 19 - Present work L. joliana Semir and Leitaõ (ine´d.) 18 36 Mansanares et al. 2002 L. mello-barretoi G.M.Barroso 19 38 Mansanares et al. 2002 L. sellowii Sch. Bip. 38 Mansanares et al. 2002 L. tomentosa (Mart. ex DC.)Sch.Bip. 17 - Coile and Jones, 1981 19 38 Mansanares et al. 2002 Lychnophora sect. Lychnophoriopsis L. heterotheca (Sch.Bip.) Coile and Jones (=L. candelabrum Sch. Bip.) 17 Coile and Jones, 1981 L. candelabrum Sch. Bip. - 36 Mansanares et al. 2002 Eremanthus E. eleagnus Sch.Bip. 15 - Turner et al. 1979 E. reflexo auriculatus Barroso (L reflexoauriculata)36Jones 1982 Minasia M. alpestris 17 - Dematteis 1998 24 mansanares, forni martins and semir numbers were obtained in the section Lychno- All analysed species showed metacentric and phora, but in the other analysed sections in this submetacentric chromosomes, although in differ- study 2n=36 was not found. Of the twenty-five ent proportions according to karyological for- species that constitute the section Lychnophorio- mula (Table 3). Similarities in karyological sym- ides (Semir 1991) only three were analysed, L. an- metry were observed in TF% values, all about 42 gelae, L. leucodendron and L. sincephala, for the (Table 3). time being appearing constant (2n=34). One spe- In Lychnophora villosissima (population 01) cies of the section Lychnocephaliopsis and one of chromosome numbers of 2n=36, 2n=37 and Chronopappus presented 2n=38. 2n=38 were observed (Figure 2-E). These num- There were some differences in chromosome bers occurred in cells from the same root-tip: length and morphology among Lycnhophora three root-tips had only 2n=36 chromosomes, one markgravii, L. rosmarinifolia and L. uniflora (Fig- had only 2n=37 and one only 2n=38 while most ure 1, Table 3). Chromosome length varied from root-tips showed all three chromosome numbers 1.10µmto2,58µm, (Table 3) and the total chro- in the same root-tip. mosome length (TCL) was relatively similar between L. rosmarinifolia and L. markgravii,itwas DISCUSSION larger in L. uniflora, although this species showed an equal or smaller chromosome number than the Among the eighteen species of Lychnophora other species here analysed. studied in this work, fourteen had their chromo-

Fig. 1 — Ideograms of Lychnophora species A: L. markgravii, 2n=38, 10m+9sm. B: L. rosmarinifolia (Pop01), 2n=36, 15m+3sm. C: L. uniflora, 2n=36, 12m+6sm. (m = metacentric; sm = submetacentric).

Table3—Chromosome numbers, karyotypical formula, chromosome length variation into species, TCL (total chromosome length), TF% (karyotipical symmetry) of Lychnophora species.

Karyotipical Length Variation TCL Species 2n TF % Formula (µm) (µm) L. markgravii 38 20m+18sm 1.14-2.23 61.50 41.26 L. rosmarinifolia 36 30m+6sm 1.10-2.10 54.88 42.34 L. uniflora 36 24m+12sm 1.30-2.58 68.69 41.35 cytotaxonomy of lychnophora 25

Fig. 2 — Mitotic metaphasis of Lychnophora sect. Lychnophora species: A − L. cryptomerioides, 2n=36 B − L. gran- mogolense, 2n=34; C − L. mutica, 2n=38; D − L. rosmarinifolia, 2n=36; E − L. villosissima, 2n=36; F-L. uniflora, 2n=36. Meiotic plates of Lychnophora sect. Lychnophora species: G − L. ramosissima, n=18; H − L. sobolifera, n=18. Bar=10µm. some number reported for the first time (Table 2, 1982; Dematteis 1998) and another Lychnophora Figures 2 and 3). The chromosome numbers ob- species (Coile and Jones 1981; Mansanares et tained (2n=34, 2n=36 and 2n=38) coincided with al. 2002). other genera of subtribe Lychnophorinae, as Er- Of currently species with cytological reports, emanthus and Minasia (Turner et al. 1979; Jones eleven show n=17 (2n=34), thirteen n=18 (2n=36) 26 mansanares, forni martins and semir

Fig. 3 — Mitotic metaphasis of Lychnophora sect. Lychnophorioides species: A − L. leucodendron, 2n=34; B − L. syn- cephala, 2n=34. Meiotic plates of Lychnophora sect. Lychnophorioides species: C − L. angelae, n=17. Mitotic metaphasis of Lychnophora sect. Chronopappus species: D-L. markgravii, 2n=38. Bar = 10µm. and eight n=19 (2n=38) (Coile and Jones 1981; shows a bigger TCL value (68.68µm) in compari- Jones 1982; Carr et al. 1999; Mansanares et al. son to L. markgravii that shows 2n=38 and 2002). TCL=61.50µm. In Lychnophora candelabrum, Coile and Jones Although all species present metacentric and (1981) reported n=17. In the present work 2n=36 submetacentric chromosomes, there is a predomi- (n=18) was observed, coinciding with the previ- nance of metacentric ones, in agreement to litera- ous report by Mansanares et al. (2002) where the ture data for the tribe Vernonieae (Ruas et al. differences in both counts were discussed. 1991; Dematteis 1996; 1998; 2002; Dematteis The chromosome number obtained for L. and Ferna´ndez 1998; 2000). granmogolensis in this study was 2n=34, in agree- Although chromosome number observed for ment with other counts for the genus L. villosissima from populations 01 and 02 was (Mansanares et al. 2002). According to constant (2n=36), individuals from population 01 Mansanares et al. (2002), the chromosome showed some variation of the number (2n=36, number reported by Carr et al. (1999) of 2n=37 and 2n-38). This is the first report for chro- 2n=18+B as L. phylicifolia is atypical for the whole mosome numerical variation in a species not re- group. In addition, L. phylicifolia was considered lated with polyploidy for Lychnophora. The char- as an incorrect identification of L. granmogolensis. acteristic diploid number of the species is 2n=36, The species Lychnophora mutica (2n=38), L. whereas 2n=37 and 2n=38 could indicate aneu- prostrata (2n=34) and L. salicifolia (2n=36) were ploidy/disploidy. analysed by Mansanares et al. (2002), chromo- Aneuploidy and disploidy occur by loss or some number counts agree with those found in gain of one or few chromosomes. In disploidy, the the present work. genotipic DNA quantities is not alterated, as oc- Karyotypic differences among three analysed curs in aneuploidy. Aneuploidy/disploidy arises species, L. markgravii, L. rosmarinifolia and L. from meiotic or mitotic deviations, or radiation uniflora, were related to number, length and chro- and chemical treatment response (Stebbins 1971; mosome morphology. The individual chromo- Malallah et al. 2001), and probably played an some length and total chromosome length (TCL) important role in the speciation mechanism of were different among the three species. Lychno- Vernonieae (Stebbins 1971; Dematteis 1996; phora uniflora (Lychnophora section) with 2n=36 1998; 2002; Mansanares et al. 2002). cytotaxonomy of lychnophora 27

Semir (1991) proposed six sections in genus by Coile and Jones (1981) as Haplostephium pas- Lychnophora. Three of them (Lychnophora, Lych- serina, can also be differentiated by their chromo- nophoriopsis and Lychnocephaliopsis) were ana- some numbers, in agreement to taxonomic posi- lysed by Mansanares et al. (2002). Here two tion established by Semir (1991). This author more sections were studied (Lychnophorioides considered these two species in Lychnophora and Chronopappus) (Table 2). based on morphological and taxonomic consid- Chromosome numbers had been registered for erations. The chromosome number here observed twenty-two, among twenty-five species of section for L. ramosissima (2n=36) is distinct of the one Lychnophora. This section grouped most of the obtained for L. passerina (2n=34) by Mansanares species with 2n=34. Lychnophora granmogolensis et al. (2002). and L. prostrata were here confirmed. Besides, Based on data presented on this and on litera- other chromosome numbers were obtained for ture related, we suggest to subtribe Lychnophori- the section, as 2n=36, observed in most of species nae four basic chromosome numbersx=15,17, and previously reported for only three species, L. 18 and 19, and the genus Lychnophora was repre- gardneri, L. pohlii and L. salicifolia, and to nine sented byx=17,18 and 19 (Mansanares et al. more in this study, L. cryptomerioides, L. itacambi- 2002). rensis, L. martiana, L. nanuzae, L. ramosissima, L. The mechanism that gives rise to chromo- rosmarinifolia, L. sobolifera, L. villosissima and L. some variations in the genus Lychnophora may be uniflora. The number 2n=38 (n=19) was earlier aneuploidy or disploidy, in agreement with Mansanares reported for two species, L. pseudovillosissima et al. (2002) and other authors who Ruas and L. mutica (Mansanares et al. 2002). studied the genera of tribe Vernonieae ( et Dematteis ; ; ; Demat- Two chromosome numbers (2n=36 and al. 1991; 1996 1998 2002 teis Ferna´ndez ; ) 2n=38) were reported for five species of section and 1998 2000 . The karyotypi- Lychnocephaliopsis (Mansanares et al. 2002). For cal evolution model in Vernonieae species sug- L. grazielae 2n=38 was observed, coinciding with gest that chromosome data from Old World species of Vernonieae are based on x=9 and 10, the number of most other species previously re- Jones ; ported by Mansanares et al. (2002). while Neotropical species on x=17 ( 1974 1979; Mathew and Mathew 1976; Turner et Section Chronopappus is constituted by two al. 1979; Ruas et al. 1991; Dematteis 1996; species, and only L. markgravii was here analysed. 1998; 2002; Dematteis and Ferna´ndez 1998; Karyotypic data obtained in this study is interest- 2000). ing because L. markgravii is more distinct in several features of their karyotype than the other two species, which are representatives of section Lych- Acknowledgements — The authors thanks to Semir IBAMA and IEF-MG for authoriztion for collect of bo- nophora. According to (1991), species of tanical material. This study represents a portion of the section Chronopappus differ from others by sev- PhD tesis of Mariana Esteves Mansanares at Universi- eral leaf characters and inflorescence. dade Estadual de Campinas. The authors thanks for the The three different chromosome numbers ob- financial support provided by FAPESP and CNPq. served occur distributed among the species, in at least four of six sections of Lychnophora sensu Semir (1991), and for the time being, is an impor- REFERENCE tant cytotaxonomic character to be used in dis- tinction of species but not in the segregation in Carr G.D., King R.M., Powell A.M. Robinson Robinson and distinct genera as suggested by (1983; H., 1999 — Chromosome numbers in Compositae, 1999), MacLeish (1984) and Hind (2000). XVIII. American Journal of Botany, 86(7): 1003- By on the one hand, the chromosome numeri- 1013. cal variations do not allow to characterize or dis- Coile N.C. and Jones Jr. S.B., 1981 — Lychnophora tinguish sections of Lychnophora,onthe other (Compositae: Vernonieae), a endemic genus to the hand, Mansanares et al. (2002) suggested that Brazilian planalto. Brittonia, 33: 528-542. Dematteis M some species could be differentiated through ., 1996 — Estudios cromosomicos en espe- comparison of this character, for example among cies Argentinas de Vernonia (Asteraceae). Bonplan- dia, 9(1-2): 103-110. the species L. ericoides and L. gardnerii, which Dematteis M., 1998 — Chromosome studies of some could be separated by their chromosome number Vernonia species (Asteraceae). Genetics and Mo- (2n=34 and 2n=36, respectively). Other species, lecular Biology, 21: 381-385. one studied in present work, Lychnophora passe- Dematteis M., 2002 — Cytotaxonomic analysis of rina and L. ramosissima, which were considered South American species of Vernonia (Vernonieae: 28 mansanares, forni martins and semir

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